Parasitoid and inquiline attack in the galls of four alien, cynipid gall wasps: host switches and the effect on parasitoid sex ratios

1. Four alien cynipid gall wasps of the genus Andricus are established and still spreading in the British Isles. The order, according to the northerliness of their distribution boundary, is: A. corruptrix→A. quercuscalicis→A. lignicola→A. kollari. All four aliens have a sexual generation in spring on Quercus cerris (introduced to Britain) and an agamic generation in autumn on native oak species. 2. For 2 years 1994 and 1995, galls of both generations of the four alien species were sampled at eight sites from the south of England to the north of Scotland to determine the parasitoid and inquiline species that attack the new galls. The spring generations of the invading species shared a parasitoid complex of four pteromalid species. Five species of inquilines and 11 species of parasitoids emerged from the autumn galls. 3. Two colonisation events were recorded for A. lignicola and A. corruptrix. On both occasions, the spring generations were found first at the new sites, indicating that the agamic generation provides the colonisers for these invading species. After colonisation, the galls of both species were attacked by parasitoids in their first season. 4. In spring, the invading species were among the most abundant cynipids at all eight sites. By sampling the whole local community of cynipid galls, it was found that the parasitoid species attacking the spring galls of the invaders seemed to have shifted their attack to the new hosts. 5. The secondary sex ratios of the parasitoid species emerging from the sexual galls of A. quercuscalicis (the smallest of the four) showed a strong and significant male bias at all sites and in both years. Parasitoid emergence from the galls of the sexual generations of the other three species (all about equal in size) was between 60 and 70% male, and variable among sites and between years.     

Survivorship of the sexual and agamic generations of Andricus quercuscalicis on Quercus cerris and Q.robur

ABSTRACT.

• 1 Survivorship within the galls of the sexual and agamic generations of Andricus quercuscalicis (Burgsdorf) is estimated from field populations.
• 2 The sexual generation on Turkey oak suffers a 20% death rate through parasitism by Mesopolobus spp. The emerging parasitoids are virtually all males.
• 3 The agamic generation on the acorn cups of English oak is not attacked by any parasites or inquilines at our Berkshire site despite the fact that most of the species which parasitize the gall in continental Europe are on the British list.
• 4 Agamic galls cause average losses of between 30% and 50% of the total acorn crop in different years. Individual trees differ greatly in their rates of loss, many losing over 90%, others losing virtually none.

     

Alien herbivores and native parasitoids: rapid developments and structure of the parasitoid and inquiline complex in an invading gall wasp Andricus quercuscalicis (Hymenoptera: Cynipidae)

Abstract.

• 1 Rapid and substantial changes have occurred in the parasitoid and inquiline community associated with the agamic galls of Andricus quercuscalicis since it invaded Britain in the late 1950s. The number of parasitoid and inquiline species has risen from one to thirteen over a 15-year period. Although the number of species has been relatively consistent over the last 8 years, the species composition has changed considerably and in a highly characteristic way during this period.
• 2 The parasitoid complex can be divided into two broadly distinct sets of parasitoid species; one set attacks only the gall former whereas the other set concentrates on the inquilines living in the wall of the gall.
• 3 The most dramatic change, however, is in the abundance of inquilines which were reported to be virtually absent in earlier studies on this community in Britain. Over a period of only 5 years, between 1988 and 1993, inquiline attack rose from less than 0.01 to an average of 0.26 inquilines per gall. The intensity of inquiline attack is geographically heterogenous, with high inquiline numbers restricted to south-east England. Because of the relatively high specificity of the parasitoids, high inquiline abundance is positively correlated with parasitoid species richness in knopper galls.
• 4 Parasitism rates, particularly on the gall former, were generally low (<10%). Over the last 5 years, however, seven parasitoid species have been consistently recorded and the mortality caused by these species has increased continuously. The species composition of the community associated with this alien gall wasp in Britain has quickly converged to the community known from its native range in continental Europe. Parasitoid species known to attack the galls of A.quercuscalisis on the continent have been recorded from it in Britain for the first time mainly in areas where inquilines have recently become abundant.
• 5 Since rates of parasitism of the gall former are still low, parasitoids are unlikely to play a major role in the population dynamics of this invading gall wasp at present, but the rapidly increasing inquiline and parasitoid attack could be a source of increased mortality for native cynipid species which are the alternative hosts of those parasitoid species.

     

Partitioning of herbivore hosts across time and food plants promotes diversification in the Megastigmus dorsalis oak gall parasitoid complex

Communities of insect herbivores and their natural enemies are rich and ecologically crucial components of terrestrial biodiversity. Understanding the processes that promote their origin and maintenance is thus of considerable interest. One major proposed mechanism is ecological speciation through host‐associated differentiation (HAD), the divergence of a polyphagous species first into ecological host races and eventually into more specialized daughter species. The rich chalcid parasitoid communities attacking cynipid oak gall wasp hosts are structured by multiple host traits, including food plant taxon, host gall phenology, and gall structure. Here, we ask whether the same traits structure genetic diversity within supposedly generalist parasitoid morphospecies. We use mitochondrial DNA sequences and microsatellite genotypes to quantify HAD for Megastigmus (Bootanomyia) dorsalis, a complex of two apparently generalist cryptic parasitoid species attacking oak galls. Ancient Balkan refugial populations showed phenological separation between the cryptic species, one primarily attacking spring galls, and the other mainly attacking autumn galls. The spring species also contained host races specializing on galls developing on different host‐plant lineages (sections Cerris vs. Quercus) within the oak genus Quercus. These results indicate more significant host‐associated structuring within oak gall parasitoid communities than previously thought and support ecological theory predicting the evolution of specialist lineages within generalist parasitoids. In contrast, UK populations of the autumn cryptic species associated with both native and recently invading oak gall wasps showed no evidence of population differentiation, implying rapid recruitment of native parasitoid populations onto invading hosts, and hence potential for natural biological control. This is of significance given recent rapid range expansion of the economically damaging chestnut gall wasp, Dryocosmus kuriphilus, in Europe.     

Host Niches and Defensive Extended Phenotypes Structure Parasitoid Wasp Communities

Oak galls are spectacular extended phenotypes of gallwasp genes in host oak tissues and have evolved complex morphologies that serve, in part, to exclude parasitoid natural enemies.Parasitoids and their insect herbivore hosts have coevolved to produce diverse communities comprising about a third of all animal species. The factors structuring these communities, however, remain poorly understood. An emerging theme in community ecology is the need to consider the effects of host traits, shaped by both natural selection and phylogenetic history, on associated communities of natural enemies. Here we examine the impact of host traits and phylogenetic relatedness on 48 ecologically closed and species-rich communities of parasitoids attacking gall-inducing wasps on oaks. Gallwasps induce the development of spectacular and structurally complex galls whose species- and generation-specific morphologies are the extended phenotypes of gallwasp genes. All the associated natural enemies attack their concealed hosts through gall tissues, and several structural gall traits have been shown to enhance defence against parasitoid attack. Here we explore the significance of these and other host traits in predicting variation in parasitoid community structure across gallwasp species. In particular, we test the “Enemy Hypothesis,” which predicts that galls with similar morphology will exclude similar sets of parasitoids and therefore have similar parasitoid communities. Having controlled for phylogenetic patterning in host traits and communities, we found significant correlations between parasitoid community structure and several gall structural traits (toughness, hairiness, stickiness), supporting the Enemy Hypothesis. Parasitoid community structure was also consistently predicted by components of the hosts'' spatiotemporal niche, particularly host oak taxonomy and gall location (e.g., leaf versus bud versus seed). The combined explanatory power of structural and spatiotemporal traits on community structure can be high, reaching 62% in one analysis. The observed patterns derive mainly from partial niche specialisation of highly generalist parasitoids with broad host ranges (>20 hosts), rather than strict separation of enemies with narrower host ranges, and so may contribute to maintenance of the richness of generalist parasitoids in gallwasp communities. Though evolutionary escape from parasitoids might most effectively be achieved via changes in host oak taxon, extreme conservatism in this trait for gallwasps suggests that selection is more likely to have acted on gall morphology and location. Any escape from parasitoids associated with evolutionary shifts in these traits has probably only been transient, however, due to subsequent recruitment of parasitoid species already attacking other host galls with similar trait combinations.     

Invaders on the move: parasitism in the sexual galls of four alien gall wasps in Britain (Hymenoptera: Cynipidae)

Four alien, host-alternating cynipid gall wasps are established in the British Isles: Andricus kollari, A. lignicola, A. quercuscalicis and A. corruptrix. Their current distributions divide Britain into four zones; all four invaders are present in the south and south-east of Britain, whereas only A. kollari is present in the far north of Scotland. The rank order, according to how far north the distributions of the four invading cynipid species reach, is: A. corruptrix→ A. quercuscalicis→ A. lignicola→ A. kollari. The life cycles of all four cynipid species involve a sexual generation in spring on Quercus cerris, and an agamic generation in autumn on Q. robur. Here we studied the parasitoid attack by four pteromalid species on the sexual generations of the invaders. We collected and reared the galls of all four species at eight sites from the south to the north of the country (two sites per zone). The geographical locations of the sites reflect the invasion history and thus the residence time of the alien species in the four zones. At each site we measured the densities of all host galls on Q. cerris and of both obligatory host-tree species. We also took a series of measures, such as host-tree density and mean host-tree size, to further characterize the tree stands. These measures are referred to as local parameters. Host densities varied between sites and between years. In A. kollari, galling rates were highest in the middle of the country (zone 2) in 1994, whereas in 1995 they increased from the south to the north. In A. lignicola, galling rates in both years were lowest at the sites in zone 3 (closest to its distribution boundary). In A. quercuscalicis, galling rates were found to be lowest at the site most to the north-west in both years, again the one furthest away from the area where this species was first recorded. Mortality caused by parasitoid attack differed from less than 10% to as high as 70% and varied between host species, sites and years. In four out of six cases the historical/regional variables (north/south and east/west) correlated significantly with parasitoid attack rates that were characteristically lowest at sites close to the distribution boundaries. Of the local factors, we found parasitoid attack rates correlated negatively in one case with host density, whereas they correlated positively in four cases with the density of alternative hosts of the parasitoids. In one of the models the local density of Q. cerris trees correlated negatively with parasitoid attack on A. quercuscalicis. For all three host species the terms retained in the minimal adequate models obtained for 1994 and 1995 differed, which might indicate that these communities of native parasitoids and invading host have not yet settled in any definite structure.     

Differential rates of invasion in three related alien oak gall wasps (Cynipidae: Hymenoptera)

Abstract. Three related species of oak gall wasps, Andricus corruptrix (Schlechtendal), A. kollari (Hartig) and A. lignicola (Hartig) have entered Britain since the introduction of Turkey oak, Quercus cerris L. in 1735. Their lifecycles involve alternating generations between an agamic generation on the native oak species (Q. petraea, Q. robur and their hybrid Q. x. rosacea), and a smaller, sexual generation on the alien Q. cerris. In examining the distributions of these insects and Q. cerris, we hypothesized that: (1) the invasion will spread more rapidly in places where both host trees are equally abundant than through regions where one of the tree species is substantially less common than the other; (2) interspecific competition between these bud‐galling species will lead to a negative correlation between their abundances at a particular site; (3) differential recruitment of natural enemies from the native hymenopteran fauna will slow the rate of spread in a species‐specific manner. A. kollari arrived nearly 200 years ago and is now found throughout the British Isles, wherever Turkey oak is grown. A. lignicola and A. corruptrix have been here for 30 years, after establishing in S.E. England. A. lignicola is in its final rapid stages of range expansion across England, southern Scotland and N.E. Scotland. A. corruptrix is just beginning to spread through Central and S.W. England. It has occupied proportionally fewer sites behind its invasion front than have the other two species, but is no less abundant at these sites. Nevertheless, distance leaps of up to 50 km were identified in A. lignicola in N.E. Scotland, and the possibility of long‐distance transport of infected trees through the horticulture and forestry trades remains. The co‐occurrence of mature individuals of both host Quercus species does appear to have increased their rates of colonization in A. lignicola and A. corruptrix. There is no evidence, however, to suggest that interspecific competition between the three alien gall formers is an important factor in determining their distributions and abundance within their invaded ranges. All three species have recruited parasitoids and inquilines rapidly from the native fauna; attack rates were highly variable, but showed no evidence of density dependence across sites.     

Honeydew feeding increased the longevity of two egg parasitoids of the pine processionary moth

The longevity of a generalist (Ooencyrtus pityocampae) and a specialist (Baryscapus servadeii) egg parasitoid of the pine processionary moth (Thau‐metopoea pityocampa) was compared under laboratory feeding conditions including water and honeydew from aphid species growing on maritime pine (Pinus pinaster) or pedunculate oak (Quercus robur). The longevity of both parasitoid species increased when specimens were fed with honeydew. This increase was larger for the generalist (3.7, 32.0 and 38.0 days) than for the specialist (3.0, 23.3 and 21.5 days) parasitoid species when fed with water, oak and pine aphid honeydew respectively. The phenology of the specialist species B. servadeii is well‐adapted with its host availability with or without food supply. The generalist species O. pityocampae could overlap its host emergence curve during 14.0–20.0 days when fed with oak and pine aphid honeydew respectively, vs. no overlap when no supplementary food was provided. Analysis of honeydew composition indicated that sugars and amino acids may have distinct effects on parasitoid longevity.     

Host–enemy interactions provide limited biotic resistance for a range-expanding species via reduced apparent competition

Aim

As species' ranges shift poleward in response to anthropogenic change, they may lose antagonistic interactions if they move into less diverse communities, fail to interact with novel populations or species effectively, or if ancestral interacting populations or species fail to shift synchronously. We leveraged a poleward range expansion in a tractable insect host–enemy community to uncover mechanisms by which altered antagonistic interactions between native and recipient communities contributed to ‘high niche opportunities’ (limited biotic resistance) for a range-expanding insect.

Location

North America, Pacific Northwest.

Methods

We created quantitative insect host–enemy interaction networks by sampling oak gall wasps on 400 trees of a dominant oak species in the native and expanded range of a range-expanding gall wasp species. We compared host–enemy network structure between regions. We measured traits (phenology, morphology) of galls and interacting parasitoids, predicting greater trait divergence in the expanded range. We measured function relating to host control and explored if altered interactions and traits contributed to reduced function, or biotic resistance.

Results

Interaction networks had fewer species in the expanded range and lower complementarity of parasitoid assemblages among host species. While networks were more generalized, interactions with the range-expanding species were more specialized in the expanded range. Specialist enemies effectively tracked the range-expanding host, and there was reduced apparent competition with co-occurring hosts by shared generalist enemies. Phenological divergence of enemy assemblages interacting with the range-expanding and co-occurring hosts was greater in the expanded range, potentially contributing to weak apparent competition. Biotic resistance was lower in the expanded range, where fewer parasitoids emerged from galls of the range-expanding host.

Main Conclusions

Changes in interactions with generalist enemies created high niche opportunities, and limited biotic resistance, suggesting weak apparent competition may be a mechanism of enemy release for range-expanding insects embedded within generalist enemy networks.     

A native and an introduced parasitoid utilize an exotic gall-maker host

Dryocosmus kuriphilus (Hymenoptera: Cynipidae) is non-native to North America and induces formation of galls on petioles and leaves of all chestnut (Castanea spp., Fagales: Fagaceae). We investigated the interactions between the gall wasp D. kuriphilus, a native parasitoid, Ormyrus labotus (Hymenoptera: Ormyridae), and a non-native parasitoid, Torymus sinensis (Hymenoptera: Torymidae). Galls were collected monthly from May to August and in January from four locations in the United States consisting of orchard-grown hybrid chestnuts (Hiram, OH and Meadowview, VA), suburban-grown ornamental Chinese chestnuts (C. mollissima) (Broadview Heights, OH), or forest-grown American chestnuts (C. dentata) (Bowling Green, KY). Parasitoids were removed from galls and T. sinensis and O. labotus were identified using PCR-markers. The relative abundance of each parasitoid was compared in relation to collection date, habitat, presence of alternative hosts, and gall characteristics. T. sinensis was collected from each location and date, and was dominant in the orchard and suburban locations. However, relatively more O. labotus were collected within the forest, which had significant oak component and alternative cynipid hosts. O. labotus was only collected in spring and early summer, indicating the use of different summer and winter hosts. Observations suggest that in addition to parasitizing D. kuriphilus, O. labotus hyperparasitizes T. sinensis. T. sinensis has a longer ovipositor than O. labotus, and parasitized larger galls. This study improves our understanding of interactions between an invasive gall wasp, an introduced parasitoid, and native parasitoids, and illustrates novel relationships that may form as exotic species expand their geographic range.     

Organization of a parasitoid community associated with a complex of galls on Atriplex spp. in southern California

Abstract. 1. Atriplex canescens (Pursh) Nuttall and A.polycarpa (Torrey) Watson (Chenopodiaceae) support twelve morphologically distinct gall types in southern California. Thirty-seven common species of parasitoids, predators and inquilines are associated with these galls. 2. The galls incited by eight members of the Asphondylia atriplicis Cockerell (Diptera: Cecidomyiidae) species complex are linked into a single, interacting community through shared hymenopterous parasitoids and inquilines. 3. Cluster analysis (UPGMA) grouped the fifteen most common species of Chalcidoidea into three host guilds of five species each: (1) specialists in tumour stem and blister leaf galls on A.canescens, (2) specialists in woolly stem galls on A.poiycarpa, and (3) generalists that attack all galls. Guild 1 dominated the galls with which it was primarily associated, while guild 3 dominated the remainder. 4. The abundances of the parasitoids of the tumour stem and blister leaf galls were negatively correlated with the abundances of two organizer species, a gall-forming inquiline, Tetrastichus cecidobroter Gordh and Hawkins, and an internal, larval—pupal parasitoid, Tetrastichus sp. B. The abundances of nine of the twelve most common chalcidoids were not correlated with the abundances of all coaccurring species in six other galls. 5. Host seasonality partly determines parasitoid population dynamics and guild structure. Parasitoid dominance increased with gall duration, suggesting that parasitoid competition depends on resource stability. The two continuously available galls were dominated by their specialist guild, while all seasonal galls were dominated by generalists. The subdominant specialists of woolly stem galls may represent competitively inferior species that utilize those galls opportunistically, because of the gall's widespread distribution and 9–10 month yearly availability. 6. Sites in the Colorado Desert and chaparral that supported several gall types showed stable relative abundances of the major parasitoid species, whereas sites in the Mojave Desert that supported only woolly stem galls had unpredictable parasitoid species assemblages. 7. The competitive success of Atriplex gall parasitoids may depend primarily on voltinism (multivoltine species dominated univoltine species) and mode of feeding (phytophagous, mixed entomophagous—phytophagous and facultatively hyperparasitic species in general dominated strict primary parasitoids).     

Fatty acid composition remains stable across trophic levels in a gall wasp community

Acquiring sufficient nutrients is particularly important for insects that are unable to synthesize certain nutrient types de novo, as is the case for numerous parasitoid species that do not synthesize lipids. The lipid reserves of parasitoids are acquired from a single host during larval development. This imposes constraints on the quantity and quality of available lipids. In the present study, the lipid dynamics throughout the trophic cascade are investigated by measuring lipogenic ability, modifications in fatty acid composition and host exploitation efficiency in species at different trophic positions within the community of parasitoids associated with the gall wasp Diplolepis rosae L. (Hymenoptera: Cynipidae). The results obtained show that lipid levels remain stable or decline after feeding in all species, indicating that none of the wasps synthesize lipids. Fatty acid composition is highly similar between the gall wasp, parasitoid and hyperparasitoid species, with the exception of the parasitoid Orthopelma mediator Thunberg (Hymenoptera: Ichneumonidae). The divergence of fatty acid composition in O. mediator suggests that this species is able to modify its fatty acid composition after the consumption of host lipids. The efficiency of exploitation of host resource, in terms of dry body mass acquired, varies among the species (41–70%), although it is high overall compared with the efficiencies reported in other animals. Hence, for parasitoid wasps that lack lipid synthesis capabilities, the efficiency of host exploitation is high and fatty acids are consumed directly from the host without modification, leading to stable fatty acid compositions throughout the trophic cascade.     

The parasitoid community of <Emphasis Type="Italic">Andricus quercuscalifornicus</Emphasis> and its association with gall size,phenology, and location

Plant galls are preyed upon by a diverse group of parasitoids and inquilines, which utilize the gall, often at the cost of the gall inducer. This community of insects has been poorly described for most cynipid-induced galls on oaks in North America, despite the diversity of these galls. This study describes the natural history of a common oak apple gall (Andricus quercuscalifornicus [Cynipidae]) and its parasitoid and inquiline community. We surveyed the abundance and phenology of members of the insect community emerging from 1234 oak apple galls collected in California’s Central Valley and found that composition of the insect community varied with galls of different size, phenology, and location. The gall maker, A. quercuscalifornicus, most often reached maturity in larger galls that developed later in the season. The parasitoid Torymus californicus [Torymidae] was associated with smaller galls, and galls that developed late in the summer. The most common parasitoid, Baryscapus gigas [Eulophidae], was more abundant in galls that developed late in the summer, though the percentage of galls attacked remained constant throughout the season. A lepidopteran inquiline of the gall (Cydia latiferreana [Tortricidae] and its hymenopteran parasitoid (Bassus nucicola [Braconidae]) were associated with galls that developed early in the summer. Parasitoids and inquilines, in general, had a longer emergence period and diapause within the gall than the gall-inducer. The association of different parasite species with galls of different size and phenology suggests that different parasite species utilize galls with slight differences in traits.     

Interactions between invading benthivorous fish and native whitefish in subarctic lakes

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The chemical composition of plant galls: are levels of nutrients and secondary compounds controlled by the gall-former?

The chemical composition of galled and ungalled plant tissue was compared in a series of experiments. Gall and adjacent plant tissue was analysed for 20 species of gall-former on 11 different plant species. There were clear differences between galled and ungalled tissue in levels of nutrients and secondary compounds. Gall tissue generally contained lower levels of nitrogen and higher levels of phenolic compounds than ungalled plant tissue. The gall tissue produced by the same plant in response to different species of gall-former differed in chemical composition, as did the gall-tissue from young and mature galls of the same species. The chemical differences between gall and plant tissues were studied in more detail in two field manipulations. Firstly, the seasonal changes in phenolic biosynthesis in Pontania proxima and P. pedunculi (Hymenoptera: Tenthredinidae) gall tissue were compared to those of their host plants, Salix alba and S. caprea. In both types of gall tissue, phenolic levels declined as the season progressed, but levels in the surrounding plant tissue increased. When the gall insects were killed with insecticide, phenolic levels in the galled tissue dropped to the same level as those in adjacent plant tissue. Secondly, the density of Cynips divisa (Hymenoptera: Cynipidae) galls on Quercus robur leaves was reduced by removing half the galls present, either those from the central region of the leaf or those from the edge. Decreasing gall density increased the size of the remaining galls and the weight of the insects, but these effects were most marked when the galls remaining were growing centrally on the leaf, i.e. when the galls from the edge had been removed. Decreasing gall density increased the nitrogen content of the remaining galls, again to a greater extent in galls growing centrally on the leaf. The results of these studies suggest that the levels of nutrients and secondary compounds in gall tissue are usually markedly different to those of surrounding plant tissue, and that gall-formers may produce species-specific and temporally variable changes in the chemical composition of gall tissue. Received: 7 July 1997 / Accepted: 29 September 1997     

The protein content of tissues in cynipid galls (Hymenoptera: Cynipidae): Similarities between cynipid galls and seeds

Cynipid galls are examples of induced plant development, where the gall inducer is in control of cell differentiation and morphogenesis of a new plant organ. This study concentrates on the tissues of the larval chamber common to all cynipid galls. The protein content of the inner gall tissue was compared to that of non‐gall plant tissues. We investigated three oak and two rose galls and their respective host plants. Total protein signatures of inner gall tissues were different from those of non‐gall plant tissues, and among the five galls. N‐terminal sequences were obtained for two abundant proteins from the inner gall tissues of D. spinosa and A. quercuscalicis, which were common to all galls, at 62 and 43 kDa. Database queries suggest the 62 kDa protein to be homologous to a protein disulphide isomerase (PDI), and the 43 kDa protein to be homologous to NAD‐dependent formate dehydrogenase (FDH). A naturally biotinylated protein was detected at 33 kDa during Western analyses with streptavidin. Western analyses revealed the presence of the biotinylated protein and PDI in the inner gall tissues of all five galls, while FDH was only detected in A. quercuscalicis and A. fecundator. PDI was also common to all non‐gall tissues, while FDH was not detected in non‐gall tissues, and the biotinylated protein was only detected in seeds. The proteins identified in the inner gall tissue suggest that (a) inner gall tissues in some galls are under respiratory stress, and (b) cynipid gall formation might involve the ectopic expression of seed‐specific proteins.     

Evolutionary shifts between host oak sections and host-plant organs in Andricus gallwasps

Abstract.— Gall-inducing insects have especially intimate interactions with their host plants and generally show great specificity with regard to both the host-plant species and the organ (e.g. flower, leaf) galled. However, the relative roles of shifts between host species and between host-plant organs in the diversification of gall-inducers are uncertain. We employ a novel and general maximum-likelihood approach to show that shifts between host-plant organs occur at a significantly greater rate than shifts between host oak sections in European Andricus gallwasps. This suggests that speciation has more often been associated with gall location shifts than with colonization of new host-plant species, and implies that it may be easier for gall-inducers to colonize new plant organs than new plant species.
Andricus gallwasps have complex life cycles, with obligate alternation of sexual and parthenogenetic generations. Our phylogenetic analyses show that a life cycle with both generations galling white oaks (section Quercus ) is ancestral, with a single shift of the sexual generation onto black oaks (section Cerris ) to generate a clade with a novel host-alternating life cycle. This new life cycle provided the opportunity for further speciation, but may have also increased the risk of extinction of one or both generations by the demographic requirement for co-existence of both host-plant groups. In summary, it appears that Andricus gallwasp radiation may be a two-level process. Speciation events often involve shifts in gall location on the same host species. However, there are only so many ways to gall an oak, and rare shifts to new oak sections may contribute greatly to long-term diversification by opening up whole new adaptive zones.     

Indirect competitive effects of stemborers on a gall community

Interspecific competition between phytophagous insects using the same host plant occurs frequently and can strongly affect population densities of competing species. Competition between gallmakers and stemborers could be especially intense because both types of herbivore are unable to avoid competition by relocation during their immature stages. For apical meristem gallmakers the main result of competition is likely to be the interruption of resources to the gall by the stemborers' devouring of stem contents. The proximate effect of such competition could be to reduce gall size, thereby increasing the number of chambers per gall unit volume, and reducing the size and potential reproductive output of the gallformer. In addition, smaller galls may be more susceptible to attack from size‐limited parasitoids, resulting in a second indirect effect of competition. Using a community of galling and stemboring insects on the saltmarsh shrub Iva frutescens L. (Asteraceae), we measured for indirect effects of competition. We examined the primary indirect effect of competition on gall midge crowding and the secondary effects on parasitism rates and parasitoid guild composition. Results indicated that galls co‐occurring with stemborers were smaller, crowding of gall inhabitants was 22% greater, and the composition of the parasitoid guild was altered relative to galls on unbored stems. The overall parasitism rate was not different between galls on bored vs. unbored stems. These results show that competition resulting from the presence of stemborers has the potential to affect the gall midge Asphondylia borrichiae Rossi & Strong (Diptera: Cecidomyiidae) and secondarily to affect its guild of hymenopteran parasitoids.     

Developmental plasticity and reduced susceptibility to natural enemies following host plant defoliation in a specialized herbivore

Host-specific phytophagous insects that are short lived and reliant on ephemeral plant tissues provide an excellent system in which to investigate the consequences of disruption in the timing of resource availability on consumer populations and their subsequent interactions with higher tropic levels. The specialist herbivore, Belonocnema treatae (Hymenoptera: Cynipidae) induces galls on only newly flushed leaves of live oak, Quercus fusiformis. In central Texas (USA) episodic defoliation of the host creates variation in the timing of resource availability and results in heterogeneous populations of B. treatae that initiate development at different times. We manipulated the timing of leaf flush in live oak via artificial defoliation to test the hypothesis that a 6- to 8-week delay in the availability of resources alters the timing of this gall former’s life cycle events, performance and survivorship on its host, and susceptibility to natural enemies. B. treatae exhibits plasticity in development time, as the interval from egg to emergence was significantly reduced when gallers oviposited into the delayed leaf flush. As a consequence, the phenologies of gall maturation and adult emergence remain synchronized in spite of variation in the timing of resource availability. Per capita gall production and gall-former performance are not significantly affected by the timing of resource availability. The timing of resource availability and natural enemies interact, however, to produce strong effects on survivorship: when exposed to natural enemies, B. treatae developing in galls initiated by delayed oviposition exhibited an order-of-magnitude increase in survivorship. Developmental plasticity allows this gall former to circumvent disruptions in resource availability, maintain synchrony of life cycle events, and results in reduced vulnerability to natural enemies following defoliation of the host plant.     

Field collection of egg parasitoids of Pentatomidae and Scutelleridae in Northwest Italy and their efficacy in parasitizing Halyomorpha halys under laboratory conditions

The invasion of Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) has caused severe economic damage in crops in North America and Europe, motivating research to identify its natural enemies, both in native and invaded areas. In its Asian native range, the main natural enemies are egg parasitoids, among which the most effective are Trissolcus japonicus (Ashmead) and Trissolcus mitsukurii (Ashmead) (Hymenoptera: Scelionidae) in China and Japan, respectively. In Europe, biology, host range, and impact of most native scelionid species are not well‐known. The present study aimed to investigate (1) presence and abundance of scelionid species that parasitize native Pentatomidae and Scutelleridae eggs in Northwest Italy, and (2) their ability to develop on H. halys eggs. During 4‐year field surveys, egg masses were collected and reared until bug nymph or adult parasitoid emergence. Then, the obtained scelionid females were tested for their ability to parasitize H. halys eggs in laboratory no‐choice experiments. Egg masses of all collected bug species were parasitized, and Telenomus spp. (Hymenoptera: Scelionidae), Trissolcus belenus (Walker), and Anastatus bifasciatus (Geoffroy) (Hymenoptera: Eupelmidae) were the most common parasitoids. In the laboratory, Trissolcus kozlovi Rjachovskij was the only species to significantly produce offspring from fresh H. halys eggs, whereas all tested Trissolcus species significantly induced host egg abortion (non‐reproductive effects). This study provides knowledge of the parasitoid species associated with native bugs, and represents a starting point to investigate the intricate interactions between native and exotic parasitoids recently found in northern Italy. These egg parasitoids could potentially be effective biocontrol agents of H. halys.